This PDF version does not have an ISBN or ISSN and is not therefore effectively published (Melbourne Code, Art. 29.1). The printed version, however, was effectively published on 6 June 2013. Wang, F. G., F. W. Xing, S. Y. Dong & M. Kato. 2013. Blechnaceae. Pp. 411–417 in Z. Y. Wu, P. H. Raven & D. Y. Hong, eds., Flora of China, Vol. 2–3 (Pteridophytes). Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. BLECHNACEAE 乌毛蕨科 wu mao jue ke Wang Faguo (王发国)1, Xing Fuwu (邢福武)1, Dong Shiyong (董仕勇)1; Masahiro Kato2 Plants mostly terrestrial, sometimes tree ferns (like small trees), rarely scandent. Rhizome mostly erect, or creeping or scandent, usually dictyostelic (Stenochlaena meristelic), scales brown, entire. Fronds monomorphic or dimorphic, mostly long stipitate; stipe with 2 to several vascular bundles, scaly at base; lamina pinnate, pinnatifid, or bipinnatifid, rarely simple, thickly papery to leathery, glabrous or usually with small scales; pinnae rarely articulate to rachis (Stenochlaena); veins free or anastomosing with 1 or more series of areoles, without included free veinlets. Sori elongate or continuous along vascular network or commissure on either side of midrib, indusiate, rarely exindusiate, rarely acrostichoid; indusium facing toward costa or costule; annulus longitudinal, interrupted. Spores elliptic, bilateral, monolete.
About (2–)14 genera and ca. 250 species: worldwide, but with greatest diversity in the S tropics; eight genera and 14 species (one endemic) in China. Stenochlaena has been separated as the sole genus of Stenochlaenaceae or Blechnaceae subfamily Stenochlaenoideae. Christenhusz et al. (Phy- totaxa 19: 48–49. 2011) listed seven genera of Blechnaceae but indicated that the actual number of genera was very uncertain and could be anywhere between two and nine, depending mainly on a better understanding of Blechnum. Shing Kunghsia. 1990. Stenochlaenaceae. In: Ching Renchang & Shing Kunghsia, eds., Fl. Reipubl. Popularis Sin. 3(1): 94–97; Wu Shiew- hung. 1999. Blechnaceae. In: Wu Shiew-hung, ed., Fl. Reipubl. Popularis Sin. 4(2): 192–215. 1a. Tree ferns with erect trunklike stem, up to ca. 1 m tall. 2a. Stem 1–2 cm in diam.; lamina bipinnatifid; sori indusiate, discrete ...... 4. Diploblechnum 2b. Stem 10–15 cm in diam.; lamina pinnate; sori exindusiate, covering pinna surface ...... 5. Brainea 1b. Rhizome creeping to suberect; plants not like tree ferns. 3a. Sori acrostichoid with sporangia covering entire abaxial surface of fertile pinna; pinnae articulate at junction with rachis; rhizome scales peltate, ca. 1 mm, caducous ...... 8. Stenochlaena 3b. Sori distinct, covered by indusia when young; pinnae not articulate; rhizome scales basifixed, 4 mm or more, often persistent. 4a. Sori at least partially discrete, parallel to costules and sometimes also costae and rachis. 5a. Rhizome short and erect; stipes tufted; pinnae separate, rachis terete; sori in lines parallel to costules, discrete ...... 6. Woodwardia 5b. Rhizome creeping; stipes distant; pinnae connected by narrow wing along rachis; sori along rachis and costae almost continuous, those along costules ± discrete ...... 7. Chieniopteris 4b. Sori forming continuous coenosori along costa of lateral pinnae. 6a. Plants epiphytic or epilithic; fronds distant; veins anastomosing ...... 2. Blechnidium 6b. Plants terrestrial; fronds clustered; veins free. 7a. Fronds monomorphic; pinnae broad ...... 1. Blechnum 7b. Fronds dimorphic; fertile pinnae very narrow, almost absent ...... 3. Struthiopteris 1. BLECHNUM Linnaeus, Sp. Pl. 2: 1077. 1753. 乌毛蕨属 wu mao jue shu Wang Faguo (王发国), Xing Fuwu (邢福武); Masahiro Kato Blechnopsis C. Presl. Plants terrestrial, moderate-sized or large. Rhizome usually erect or ascending, stout, dictyostelic, densely scaly; scales dark brown to brown, glossy, lanceolate, entire, basifixed. Fronds clustered, long stipitate; stipe robust, scaly at base, above naked; lamina pinnate, with reduced auricular lower pinnae, leathery; pinnae entire, linear, margin entire or serrulate; terminal pinna entire, similar to lateral pinnae; veins free, parallel, simple or forked. Sori linear, forming long coenosori adjacent and parallel to costa; indu- sium attached to commissure, facing toward costa; annulus longitudinal, interrupted, of 14–28 cells. Spores elliptic, usually smooth, with perispore. Up to 200 species including species of Struthiopteris, which is usually synonymized under Blechnum: pantropical; one species in China. Smith et al. (Taxon 55: 716. 2006) and Christenhusz et al. (Phytotaxa 19: 48–49. 2011) include the three following genera, Blechnidium, Struthiopteris, and Diploblechnum, within Blechnum.
1 South China Botanical Garden, Chinese Academy of Sciences, 723 Xingke Road, Tianhe District, Guangzhou, Guangdong 510650, People’s Republic of China. 2 Department of Botany, National Museum of Nature and Science, Amakubo 4-1-1, Tsukuba 305-0005, Japan.
411 412 BLECHNACEAE
1. Blechnum orientale Linnaeus, Sp. Pl. 2: 1077. 1753. Exposed shrubby or low hillsides; 200–1000 m. Chongqing, Fu- jian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangxi, Si- 乌毛蕨 wu mao jue chuan, Taiwan, Xizang, Yunnan, Zhejiang [Japan; tropical Asia, Aus- Blechnopsis orientalis (Linnaeus) C. Presl. tralia, Pacific islands]. Rhizome dark brown, erect, short, densely scaly; scales Reviewer Christenhusz notes that, in an unpublished molecular dark brown at center, brown near margin, narrowly linear, ca. 1 phylogeny, Blechnum orientale is more closely related to the New cm, entire. Stipe 10–60 cm, 3–10 mm in diam., base dark World genus Salpichlaena J. Smith and to Stenochlaena than Blech- brown and covered with scales as rhizome; lamina imparipin- num s.s., typified by B. occidentale Linnaeus, or Struthiopteris s.s., nate, monomorphic, ovate-lanceolate, 55–100 × 20–60 cm, sub- represented by B. spicant (Linnaeus) Roth. leathery; pinnae numerous, close, alternate; lower ones con- Plants with cristate apices to the lateral pinnae have been called tracted to small rounded auricles, 0.3–1 cm; upper ones oblique, “Blechnum orientale var. cristatum” (冠羽乌毛蕨 guan yu wu mao distant, linear or linear-lanceolate, 10–30 × 0.8–1.8 cm, base jue), but that name appears not to have been validly published. It has sessile, rounded or subtruncate, or adnate, decurrent to rachis on been attributed to J. Smith, but it is not mentioned in the publication basiscopic side, gradually narrowed to acuminate apex, terminal cited in FRPS, i.e., Seemann, Bot. Voy. Herald 10: 427. 1857 (in- pinna similar to middle pinnae; veins free, parallel, simple or correctly cited as “Bot. Beechey Voy. 1”), which refers only to B. forked near costa, close. 2n = 66. orientale.
2. BLECHNIDIUM T. Moore, Brit. Ferns 2: 210. 1860. 乌木蕨属 wu mu jue shu Wang Faguo (王发国), Xing Fuwu (邢福武); Masahiro Kato Plants epiphytic, small to moderate-sized. Rhizome creeping, dark brown, dictyostelic, densely scaly; scales rufous, lanceolate or ovate-lanceolate. Fronds distant, long stipitate; stipe grooved adaxially, sparsely scaly at base; lamina pectinately pinnatisect, lanceolate in outline, subleathery, both surfaces glabrous; pinnae adnate, oblong, margin entire, apex subacute or obtuse; lower few pairs of pinnae shorter; veins anastomosing with 1–3 rows of areoles. Sori linear, forming coenosori adjacent and parallel to costa; indusium attached to commissure, facing toward costa. Spores elliptic, with perispore. One species: China, India, Myanmar. Blechnidium is most closely related to Struthiopteris and isolated from the rest. It is included within Blechnum by Christenhusz et al. (Phytotaxa 19: 34. 2011).
1. Blechnidium melanopus (Hooker) T. Moore, Brit. Ferns 2: in diam., grooved adaxially, sparsely scaly at base; lamina 210. 1860. brownish green when dry, lanceolate, 15–25 × 3–7 cm, sub- leathery, both surfaces glabrous, gradually narrowed to base, 乌木蕨 wu mu jue pectinate, acuminate; pinnae 15–25 pairs, adnate; middle ones oblong, usually subfalcate, 1.5–5 × 0.5–1 cm, entire, subacute Blechnum melanopus Hooker, Sp. Fil. 3: 64. 1859; Blech- or obtuse; lower pinnae shortened, 0.4–1.3 cm; veins inconspic- nidium plagiogyriifrons (Hayata) Hayata; Blechnum plagio- uous, anastomosing with 1–3 rows of hexagonal areoles. Sori gyriifrons Hayata. linear, adjacent and parallel to costa; indusium attached to com- Rhizome dark brown, long creeping, ca. 3 mm in diam., missure, linear, facing toward costa. Spores elliptic, with peri- densely scaly; scales rufous, lanceolate to ovate-lanceolate, ca. spore. 5 mm, membranous, subentire, acuminate or fibriform. Fronds Epiphytic on tree trunks or epilithic in forests; 800–2800 m. distant, long stipitate; stipe dark brown, 12–25 cm, ca. 1 mm Guizhou (Guiding), Taiwan, Yunnan [India, Myanmar].
3. STRUTHIOPTERIS Scopoli, Meth. Pl. 25. 1754. 荚囊蕨属 jia nang jue shu Wang Faguo (王发国), Xing Fuwu (邢福武); Masahiro Kato Plants epilithic. Rhizome erect or ascending, short, dictyostelic, scaly; scales brown, linear or lanceolate, entire. Fronds tufted, ± dimorphic, often shortly stipitate; lamina pectinately pinnatisect, oblanceolate in outline, narrowed downward, leathery; pinnae spreading, oblong-subtriangular, base decurrent to rachis wing, margin entire; lower pinnae much shorter; veins inconspicuous, free, furcate, basal ones always trifurcate, not reaching lamina margin. Sori linear, forming coenosori adjacent and parallel to costa; indu- sium facing toward costa, thinly papery, covering young sporangia, opening when mature; spores elliptic, perispore corrugated. 2n = 62, 68.
About ten species: mainly distributed in temperate areas of the N Hemisphere and Australia; two species (one endemic) in China. Struthiopteris is included within Blechnum by Christenhusz et al. (Phytotaxa 19: 39. 2011).
BLECHNACEAE 413
1a. Sterile lamina 2–5 cm wide; stipe 3–24 cm ...... 1. S. eburnea 1b. Sterile lamina 5–8 cm wide; stipe almost absent ...... 2. S. hancockii
1. Struthiopteris eburnea (Christ) Ching, Sunyatsenia 5: 243. 1b. Struthiopteris eburnea var. obtusa (Tagawa) Tagawa, Acta 1940. Phytotax. Geobot. 14: 192. 1952.
荚囊蕨 jia nang jue 天长罗蔓蕨 tian chang luo man jue Rhizome erect and short, or ascending and long, densely Spicantopsis eburnea var. obtusa Tagawa, Acta Phytotax. scaly; scales brown, lanceolate, 5–6 mm, thickly membranous, Geobot. 9: 88. 1940. margin entire or with few teeth, apex fibriform. Fronds tufted, dimorphic; stipe stramineous, almost absent or 3–24 cm, 1–2 Lamina to 3 cm wide; pinnae obtuse or rounded. mm in diam., base scaly, glabrous upward; lamina pectinate- ● On limestone of central mountain ranges; (?800–)1200–2200 m. pinnatisect, dark green or brown when dry, linear-oblanceolate, Taiwan. 15–45 cm, 2–5(–8) cm above middle, thickly leathery, glabrous; pinnae 20–40 pairs, spreading, lower ones gradually shortened, 2. Struthiopteris hancockii (Hance) Tagawa, Acta Phytotax. basal pair small auriculate; upper pinnae oblong, 1.5–3 × ca. 0.5 Geobot. 14: 192. 1952. cm, base adnate to rachis, margin entire, slightly narrowed to 宽叶荚囊蕨 kuan ye jia nang jue apex, slightly reflexed, spreading, close or distant; veins incon- spicuous, pinnate, furcate, not reaching lamina margin; rachis Blechnum hancockii Hance, J. Bot. 21: 267. 1883; Lo- stramineous, glabrous, shallowly grooved adaxially. Fertile maria apodophylla Baker; L. hancockii (Hance) Baker; Spic- lamina as long as but narrower than sterile lamina. Sori linear, antopsis hancockii (Hance) Masamune; S. niponica (Kunze) from near costa to near laminar margin, single on either side of Nakai var. hancockii (Hance) Nakai. costa. 2n = 66*. Rhizome erect, short, densely scaly; scales dark brown, ● On rocks or limestone; 500–2200 m. Anhui, Fujian, Guangxi, linear, ca. 1.5 cm, membranous, entire, apex subulate. Fronds Guizhou, Hubei, Hunan, Sichuan, Taiwan. tufted, subdimorphic with fertile lamina narrower than sterile 1a. Lamina to 5 cm wide; pinnae acute ...... 1a. var. eburnea lamina, subsessile or shortly stipitate; lamina pectinate-pinnati- 1b. Lamina to 3 cm wide; pinnae obtuse sect, brown when dry, elliptic or oblanceolate, broadest at or or rounded ...... 1b. var. obtusa above middle, 30–40 × 5–8 cm, thickly leathery, glabrous, base gradually narrowed with small auricles, apex acuminate or 1a. Struthiopteris eburnea var. eburnea acute; pinnae 20–40 pairs, spreading, close or distant, linear- 荚囊蕨(原变种) jia nang jue (yuan bian zhong) oblong, slightly falcate, largest 2.5–4 × ca. 0.5 cm, in fertile fronds 1.5–3 × ca. 0.3 cm, base adnate to rachis, margin entire, Blechnum eburneum Christ, Bull. Acad. Int. Géogr. Bot. apex acute, slightly reflexed when dry; veins inconspicuous, 11: 233. 1902; Lomaria eburnea (Christ) Ching; Spicantopsis pinnate, furcate, ending in submarginal fusiform hydathodes; eburnea (Christ) Tagawa. rachis brown or stramineous, shallowly grooved adaxially, Lamina to 5 cm wide; pinnae acute. sparsely scaly; scales brown, lanceolate. Sori linear, single on either side of costa, occupying most of pinna lamina. ● On rocks or limestone; 500–1800 m. Anhui, Fujian, Guangxi, Guizhou, Hubei, Hunan, Sichuan. Forests. Taiwan [Japan].
4. DIPLOBLECHNUM Hayata, Bot. Mag. (Tokyo) 41: 702. 1927. 扫把蕨属 sao ba jue shu
Wang Faguo (王发国), Xing Fuwu (邢福武); Masahiro Kato
Tree ferns, terrestrial. Stem black, erect, dictyostelic, covered with stubs of persistent frond bases, apex densely scaly; scales dark brown, narrowly triangular-lanceolate, entire. Fronds tufted, shortly stipitate; stipe scaly; lamina bipinnatifid, elliptic-lanceolate in outline, narrowed downward, papery, both surfaces glabrous, apex acuminate; pinnae pinnatifid, oblong-lanceolate, base decurrent to narrow rachis wing with triangular lobes between pinnae, apex acuminate; lower pinnae much shorter, basal ones triangular auri- cles; lobes oblong or triangular-oblong, margin entire, apex acute; veins free, pinnate, secondary veins 2- or 3-furcate, not reaching lamina margin; rachis and costae grooved adaxially. Fertile lamina as large as sterile lamina, slightly narrowed, pinnae caudate. Sori linear, costal, adjacent and parallel to costules on straight unbranched fertile veins parallel to costa; indusia facing toward costa, thin, covering young sporangia, opening when mature; spore reniform.
One species: Taiwan and SE Asian and Pacific islands.
The genus Diploblechnum is an ingroup of Blechnum (R. Cranfill, Y. Nakahira & M. Kato, unpubl. data), although the two are distinct morpho- logically.
414 BLECHNACEAE
1. Diploblechnum fraseri (A. Cunningham) De Vol in H. L. Li subulate. Fronds tufted; stipe ca. 3 cm; lamina bipinnatifid, et al., Fl. Taiwan 1: 153. 1975. elliptic-lanceolate in outline, narrowed downward, up to 40 × 10–14 cm, papery, both surfaces glabrous; pinnae to 30 pairs, 扫把蕨 sao ba jue pinnatifid, oblong-lanceolate, 6.5–8.5 × 1.4–1.8 cm, bases de- Lomaria fraseri A. Cunningham, Companion Bot. Mag. 2: current with triangular lobes between, apex acuminate; lower 364. 1836; Blechnum fraseri (A. Cunningham) Luerssen; B. pinnae much shorter, basal ones triangular auricles; lobes 13–15 integripinnulum Hayata; Diploblechnum integripinnulum (Ha- pairs, oblong or triangular-oblong, up to 9 × 3 mm, margin yata) Hayata; L. fraseri var. philippinensis Christ; Struthiopteris entire, apex acute. Fertile lamina as large as sterile lamina, fraseri (A. Cunningham) Hayata; S. integripinnulum (Hayata) slightly narrowed, pinnae caudate. Sori linear, adjacent and Hayata. parallel to costules; indusia brown.
Stem black, up to ca. 0.8 m tall, 1–2 cm in diam., covered Cloud forests, damp and somewhat exposed path sides; 1200– with stubs of persistent frond bases, apex densely scaly; scales 2000 m. SE Taiwan [Indonesia (Borneo), Malaysia, Philippines; Pacific dark brown, narrowly triangular-lanceolate, ca. 1 cm, entire, islands (New Zealand)].
5. BRAINEA J. Smith, Cat. Ferns Roy. Gard. Kew, 5. 1856. 苏铁蕨属 su tie jue shu Wang Faguo (王发国), Xing Fuwu (邢福武); Masahiro Kato Bowringia Hooker (1853), not Champion ex Bentham (1852). Tree ferns, terrestrial. Stem erect, very stout, dictyostelic, woody, apex scaly; scales rufous, linear, membranous, acuminate. Fronds clustered in terminal crown, slightly dimorphic; stipe with basal part scaly; lamina pinnate, elliptic-lanceolate in outline, leathery, abaxially with some small scales along costa and veins; pinnae opposite or alternate, subsessile, entire, linear to narrowly oblong; basal pinnae slightly shorter; veins free, simple or 1- or 2-forked, except for costal row of subtriangular areoles; fertile pinnae somewhat shorter, margin sometimes irregularly lobed; rachis grooved adaxially. Sori borne on costal veins, abundant, covering whole abaxial surface of pinnae when mature, indusium absent. 2n = 66. One species: widely distributed in tropical Asia. The monotypic genus Brainea is isolated from the rest of the family.
1. Brainea insignis (Hooker) J. Smith, Cat. Ferns Roy. Gard. cm, slightly narrowed downward, leathery, abaxially with some Kew, 5. 1856. small scales along costa and veins; pinnae 30–55 pairs, opposite or alternate, subsessile, linear to narrowly oblong; middle pin- 苏铁蕨 su tie jue nae 10–15 × 0.7–1.2 cm, base asymmetrical, subauriculate, margin serrulate; basal pinnae slightly shortened; veins free, Bowringia insignis Hooker, Hooker’s J. Bot. Kew Gard. simple or 1- or 2-forked, except for costal row of areoles subtri- Misc. 5: 237. 1853; Brainea formosana Hayata; B. insignis var. angular; fertile pinnae somewhat shorter, margin sometimes formosana (Hayata) Tagawa. irregularly lobed; rachis stramineous, grooved adaxially. Sori Stem blackish brown, erect, stout, up to ca. 1 m tall, 10–15 borne on costal veins, exindusiate, abundant, covering whole cm in diam., woody, apex scaly; scales rufous, linear, up to abaxial surface of pinnae when mature. 2n = 66. 3 cm, membranous, acuminate. Fronds clustered in terminal Damp and exposed hillsides; 300–1700 m. Fujian, Guangdong, crown, slightly dimorphic; stipe brownish stramineous, 10–30 Guangxi, Guizhou (Anlong, Zhenning), Hainan, C Taiwan, Yunnan cm, basal part scaly; lamina pinnate, elliptic-lanceolate, 50–100 [tropical regions of Asia].
6. WOODWARDIA J. E. Smith, Mém. Acad. Roy. Sci. (Turin) 5: 411. 1793. 狗脊属 gou ji shu Wang Faguo (王发国), Xing Fuwu (邢福武); Masahiro Kato Plants terrestrial, of moderate to large size. Rhizome erect or ascending, sometimes creeping, stout, dictyostelic, densely scaly; scales brown, non-clathrate, basifixed. Fronds tufted, long stipitate; lamina deeply bipinnatifid, lower part often pinnate, elliptic in outline, papery or thinly leathery; pinnae pinnatifid, narrowly oblong in outline, gradually becoming narrowly triangular-ovate, sometimes stalked, glabrous, margin entire or serrulate; veins anastomosing to form a series of areoles along costae and costules, free to margin, simple or forked. Sori discrete, linear, elliptic, or crescent-shaped, occupying costular areoles, discrete, attached to outer areole-forming veins, superficial or sunken; indusia dark brown, facing toward costa, thickly papery; sporangia with long stalk, annulus of 17–24 thickened cells. Spores elliptic, perispore rugose. About ten species: temperate to tropical areas in Asia, Central and North America, and Europe; five species in China.
BLECHNACEAE 415
1a. Rachis with large subterminal bulbils on rachis ...... 1. W. unigemmata 1b. Rachis without such bulbils. 2a. Base of lower pinnae asymmetrical with 1–3 basiscopic lobes lacking; sori sunken, crescent-shaped or elliptic. 3a. Lamina 35–45 cm; base of lower pinnae lacking 1 basiscopic lobe; lobes acute or acuminate ...... 2. W. orientalis 3b. Lamina 35–120 cm; base of lower pinnae lacking 1–3 basiscopic lobes; lobes acuminate or caudate ...... 3. W. prolifera 2b. Base of lower pinnae ± symmetrical, basal pairs of lobes shortened or slightly so; sori not sunken, linear. 4a. Basal pairs of lobes shortened, apex obtuse ...... 4. W. japonica 4b. Basal pairs of lobes slightly shortened, lanceolate, apex acute ...... 5. W. magnifica
1. Woodwardia unigemmata (Makino) Nakai, Bot. Mag. pairs, close, oblique, oblong, 3–5.5(–7) × 0.8–1(–1.3) cm, usu- (Tokyo) 39: 103. 1925. ally broadest at base, margin cuspidately serrulate, apex acute or acuminate; veins obvious, anastomosing to form 1 row of 顶芽狗脊 ding ya gou ji areoles along costae and 2 or 3 rows of discrete, polygonal are- Woodwardia radicans (Linnaeus) Smith var. unigemmata oles, remainder free to margin, simple or forked. Small bulbils Makino, J. Jap. Bot. 2(2): 7. 1918; W. himalaica Ching & S. K. borne on adaxial surfaces of pinna lobes or not. Sori occupying Wu; W. latiloba Ching & P. S. Chiu; W. maxima Ching; W. costular areoles, crescent-shaped or elliptic, sunken in rimmed yunnanensis Ching & P. S. Chiu. depressions; indusia dark brown, thickly papery. 2n = 136. Rhizome decumbent, dark brown, stout, up to 3 cm in Roadsides, mountain slopes; ca. 500 m. Anhui, Fujian, Guang- diam., densely scaly; scales brown, lanceolate, 2.2–2.7 cm, dong, Guangxi, Hunan, Jiangxi, ?Taiwan, Zhejiang [Japan, Philippines]. membranous, entire, apex long acuminate. Stipes tufted, 30– 3. Woodwardia prolifera Hooker & Arnott, Bot. Beechey Voy. 100 cm, 5–8 mm in diam., base densely scaly; upper part of 275. 1838. stipe and rachis with fewer brown, fibriform scales; lamina 珠芽狗脊 deeply bipinnatifid, elliptic or ovate-lanceolate, 30–100 × 20– zhu ya gou ji 60 cm, leathery; pinnae 7–15 pairs, subsessile or shortly stalked, Woodwardia angustiloba Hance; W. exaltata Nakai; W. broadly lanceolate; middle pinnae 15–35 × 4–12 cm, cut 3/4 orientalis Swartz var. formosana Rosenstock; W. orientalis var. way to costa, base rounded-truncate, unequal, apex caudate; prolifera (Hooker & Arnott) Ching; W. prolifera var. formosana lobes 14–20 pairs, close, oblique, oblong-lanceolate, sometimes (Rosenstock) Ching. falcate, lower pairs of lobes slightly shortened, middle lobes 1– 6 × 0.8–1.2 cm, margin serrulate with sharp, long teeth; veins Rhizome decumbent, dark brown, stout, densely scaly; anastomosing with 2 or 3 rows of areoles along costae and scales red-brown, lanceolate, 2–4 cm, membranous, entire or costules, marginally free, simple or forked. Bulbils single or with a few teeth, fibriform. Stipes close, 30–110 cm, 3–15 mm multiple, large, scaly, subterminal below rachis tip at base of in diam., base densely scaly; upper part of stipe and rachis upper pinna. Sori occupying costular areoles, oblong, sunken in sparsely covered with brown, broadly lanceolate scales, surface rimmed depressions; indusia attached to outer vein, dark brown, rough with scale scars; lamina deeply bipinnatifid, brown or thickly membranous. 2n = 68. slightly green when dry, oblong-ovate or elliptic, 35–120 × 30– 40 cm, leathery, glabrous, apex acuminate; pinnae 5–9(–13) In shrubs by roadsides, forests; 400–3000 m. Fujian, Gansu, pairs, shortly stalked, lanceolate; lower and middle pinnae 10– Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Shanxi, Si- 30 × 4–9 cm, deeply pinnatifid to 1–2 mm from costa, base chuan, Taiwan, Xizang, Yunnan [Bhutan, India, Japan, Kashmir, Myan- asymmetrical with 1–3 basiscopic lobes lacking, apex long acu- mar, Nepal, Pakistan, Philippines, Vietnam]. minate or caudate; lobes 10–14(–24) pairs, close or separate, 2. Woodwardia orientalis Swartz in Schrader, J. Bot. 1800(2): oblique, oblong-lanceolate, 3–7(–9) × 0.5–0.9 cm, slightly nar- 76. 1801. rowed to base, margin cuspidately serrulate, occasionally lobed, apex long acuminate or caudate; veins obvious, anastomosing 东方狗脊 dong fang gou ji with 1 row of areoles along costae and 2 or 3 rows of discrete, Woodwardia radicans (Linnaeus) Smith var. orientalis polygonal areoles, free distally, simple or forked. Leaf-bearing (Swartz) Swartz. bulbils small and usually abundant on adaxial surfaces of pinna lobes. Sori occupying costular areoles, crescent-shaped or ellip- Rhizome decumbent, dark brown, stout, densely scaly; tic, sunken in rimmed depressions; indusia dark brown, thickly scales dark brown, lanceolate, 1–4 cm, entire, membranous, papery. 2n = 68. apex fibriform. Stipes close, 20–55 cm, 3–6 mm in diam., base densely scaly; upper part of stipe and rachis sparsely covered Mountain slopes, open and wet places in sparse forests, near with brown, broadly lanceolate scales; lamina deeply bipinnati- streams; 100–1100 m. Anhui, Fujian, Guangdong, Guangxi, Hunan, fid, brown or slightly green when dry, ovate, 35–45(–70) × 15– Jiangxi, Taiwan, Zhejiang [Japan]. 45 cm, leathery, glabrous, base rounded-truncate, apex acumi- Herbarium and field observations of Woodwardia prolifera show nate; pinnae 6–8 pairs, shortly stalked, lanceolate; lower and that there is ecological variation in the frequency of bulbils on the lami- middle pinnae 10–30 × 4–9 cm, deeply pinnatifid to 2–3(–4) na; some mature fronds are not bulbiliferous. mm from costa, base asymmetrical with 1 basiscopic lobe The name “Woodwardia radicans (Linnaeus) Smith var. prolifera lacking, apex acuminate, basal pinnae shortened; lobes 10–18 C. Christensen, Index Filic. 658. 1905” (Acta Phytotax. Sin. 12: 244.
416 BLECHNACEAE
1974; and FPRS 4(2): 201. 1999) appears to be a mistaken joining of Ridges, exposed slopes, shaded forests; 300–1500 m. Widely dis- the end of one line “= W. radicans var.” with the beginning of another, tributed south of the Chang Jiang and in Taiwan [Japan, Korea, Viet- the next species in the list, “prolifera.” nam]. 4. Woodwardia japonica (Linnaeus f.) Smith, Mém. Acad. Woodwardia japonica is variable in the length, number, and Roy. Sci. (Turin) 5: 411. 1793. arrangement of the pinnae and in the shape of the pinna lobes.
狗脊 gou ji 5. Woodwardia magnifica Ching & P. S. Chiu, Acta Phytotax. Sin. 12: 247. 1974. Blechnum japonicum Linnaeus f., Suppl. Pl. 445. 1782; 滇南狗脊 Woodwardia affinis Ching & P. S. Chiu; W. intermedia Christ; dian nan gou ji W. japonica var. contigua Ching & P. S. Chiu; W. omeiensis Rhizome decumbent, dark brown, stout, densely scaly; Ching ex P. S. Chiu. scales brown or dark brown, lanceolate, ca. 1.5 cm, membra- Rhizome decumbent, dark brown, stout, 3–5 cm in diam., nous, entire, apex fibriform. Stipes close, brown-stramineous, densely scaly; scales dark brown, lanceolate or linear-lanceo- 35–76 cm, 6–10 mm in diam., base densely scaly, sparsely late, ca. 1.5 cm, membranous, entire, apex acuminate, some- scaly upward; lamina bipinnatifid, brown or brown-green when times fibriform. Stipes close, 15–70 cm, 5–8 mm in diam., base dry, oblong-ovate, 60–95 × 30–65 cm, leathery, glabrous, apex densely scaly; upper part of stipe and rachis sparsely covered long acuminate; pinnae 13–15 pairs, alternate or lower ones with brown, fibriform scales; lamina bipinnatifid, elliptic or opposite, oblique, subsessile or shortly stalked, basal pair lanceolate, 25–85 × 18–45 cm, leathery, apex acuminate; pinnae slightly shortened; middle ones linear-lanceolate, 30–35 × 4–8 7–15 pairs, subsessile or shortly stalked, broadly lanceolate; cm, base rounded-truncate or rounded-cuneate, margin pinnati- middle pinnae 12–25 × 2–4 cm, base acroscopically truncate, fid, apex caudate-acuminate; lobes 20–22 pairs, alternate, lower basiscopically rounded, margin cut 1/2 way to costa, apex acu- ones slightly shortened; middle one longest, lanceolate or nar- minate; lobes 11–16 pairs, close, oblique, hemielliptic or hemi- rowly triangular-lanceolate, 2.6–5 × 1–1.5 cm, margin serrulate, ovate, lowest pair shortened, basiscopic lobe rounded, ovate or apex acute; veins obvious, anastomosing with 1 row of areoles auriculate, 5–10 mm, apex rounded; middle lobes 1.3–2.2 × along rachis, costae, and costules, distally simple or forked, ending in submarginal fusiform hydathodes; rachis or costae 0.7–1 cm, margin serrulate, apex acute; veins anastomosing densely covered with brown, fibriform, deciduous scales. Sori with 2 or 3 rows of areoles along costae and costules, distally occupying costular areoles, linear, discrete; indusia dark brown. free, simple or forked. Sori occupying costular areoles, discrete, linear, sunken; indusia brown. 2n = 68. Near forests, roadsides; 1400–1600 m. Yunnan [Vietnam].
7. CHIENIOPTERIS Ching, Acta Phytotax. Sin. 9: 37. 1964. 崇澍蕨属 chong shu jue shu Wang Faguo (王发国), Xing Fuwu (邢福武); Masahiro Kato Plants terrestrial, of small to moderate size. Rhizome long creeping, dark brown, apex covered with brown, lanceolate scales. Fronds distant, long stipitate; lamina simple, trifid, or deeply pinnatifid, thickly papery or subleathery, glabrous; pinnae (or lobes) 1– 5 pairs, lanceolate, base slightly narrowed, apex acuminate, terminal pinna if present similar to but longer than lateral ones; veins anastomosing with 3 or 4 rows of areoles, marginal veins free. Sori oblong or linear, borne along costal and costular areoles; indusia facing toward costa, dark brown, linear-oblong, papery. Spores elliptic, perispore rugose, granular.
Two species: China, Japan, Vietnam; two species in China.
Chieniopteris is treated as a section of the genus Woodwardia by Cranfill and Kato (in Subhash Chandra and Mrittunjai Srivastava, eds., Pteridol. New Millennium, 25–48. 2003), which was followed by Christenhusz et al. (Phytotaxa 19: 27. 2011). 1a. Lamina simple or imparipinnate with 1–4 pairs of lateral pinnae and longer terminal pinna ...... 1. C. harlandii 1b. Lamina pinnatifid, with pinnatifid apex, lateral pinnae 5–7 pairs, ± lobed ...... 2. C. kempii
1. Chieniopteris harlandii (Hooker) Ching, Acta Phytotax. cm; stipe 12–80 cm, base dark brown, scaly; lamina simple, Sin. 9: 39. 1964. ternate, or deeply pinnatifid, gray-green or brown when dry, 崇澍蕨 chong shu jue thickly papery or subleathery, glabrous; pinnae (or lobes) 1–4 pairs, opposite, 4–5 cm apart, lanceolate, base adnate to rachis Woodwardia harlandii Hooker, Fil. Exot. t. 7. 1857; and decurrent, margin entire or undulate, usually reflexed when Lorinseria harlandii (Hooker) J. Smith. dry, apex acuminate; basal pinnae 20–29 × 2–3 cm, upper pin- Rhizome long creeping, dark brown, 4–6 mm in diam., nae shortened, terminal pinna similar to but longer and broader densely scaly; scales brown, lanceolate, 4–6 mm, membranous, than lateral ones; rachis wing often very narrow or lacking be- margin entire or with few hairlike teeth. Fronds distant, subdi- tween basal pinnae; veins anastomosing with 1 row of costal morphic with fertile lobes narrower than sterile lobes, 30–120 areoles, 2 or 3 rows of hexagonal areoles, marginal veins free.
BLECHNACEAE 417
Sori linear, interrupted, 10–22 mm, borne along costal and cos- brous, base rounded-truncate, apex acuminate; sterile lamina tular areoles; indusia rufous when mature, papery. pinnatifid, basal lobes largest, lanceolate, 6.5–10 × 1–2.5 cm, base slightly narrowed, united with broad wing, margin serru- Valleys, damp forests; 400–1300 m. Fujian, Guangdong, Guangxi, Guizhou (Libo), Hainan, S Hunan, Taiwan [Japan, Vietnam]. late, apex acuminate; fertile lamina bipinnatifid; pinnae 5–7 pairs, opposite, oblique or basal pair subspreading, pinnatifid; 2. Chieniopteris kempii (Copeland) Ching, Acta Phytotax. Sin. basal pair largest, elliptic-lanceolate, 11–20 × 4.5–10 cm, base 9: 39. 1964. narrowed, sessile or shortly stalked, ca. 3 cm apart from above, 裂羽崇澍蕨 lie yu chong shu jue pinnatipartite to broad wing of costa, apex acuminate; middle lobes lanceolate or linear, 8–9 × 1.2–1.5 cm, margin irregularly Woodwardia kempii Copeland, Philipp. J. Sci., C, 3: 280. undulate or lobed, entire, or with irregular and distant serrae, 1908; W. harlandii Hooker var. takeoi (Hayata) Masamune; W. basal lobes much shortened or rounded-auriculate; veins incon- heteropinnata B. S. Wang; W. takeoi Hayata. spicuous but midvein prominent on both surfaces, anasto- Rhizome long creeping, dark brown, 4–7 mm in diam., mosing with 1 row of long, narrow areoles along costa, second- densely scaly; scales brown, lanceolate, 4–5 mm, membranous, ary veins anastomosing with 2 rows of areoles, free near mar- entire, apex acuminate. Fronds distant, subdimorphic, fertile gin. Sori linear, 5–17 mm, borne in costal, rachis, or midvein fronds larger and more deeply dissected than sterile fronds; areoles; indusium dark brown when mature, papery. stipe 30–70 cm, base dark brown, densely scaly, upward sparsely scaly and glabrous, brown-stramineous; lamina brown Forests, damp forests, along ridges; 400–800 m (in Taiwan). when dry, deltoid-ovate, 13–26 × 11–28 cm, subleathery, gla- Fujian, Guangdong, Guangxi, N Taiwan [Japan].
8. STENOCHLAENA J. Smith, J. Bot. 3: 401. 1841. 光叶藤蕨属 guang ye teng jue shu Dong Shiyong (董仕勇); Masahiro Kato Terrestrial and climbing plants. Rhizome creeping or climbing, stout, terete, meristelic, with 40 or more vascular bundles in 3 circles, scaly only at apex; scales brown to blackish, orbicular, ovate, or ovate-lanceolate, margin uneven, peltate, caducous. Fronds distant, strongly dimorphic, stipe glabrous except when very young; lamina imparipinnate, leathery or stiffly papery, both surfaces glabrous; sterile pinnae broadly lanceolate, articulate to rachis (apical pinna not articulate), shortly stalked, with single gland on base of each costa, margin sharply toothed; veins anastomosing, forming single row of areoles along each side of costa, other veins free; fertile pinnae linear, lamina very reduced, margin entire. Sporangia acrostichoid, covering whole abaxial surface of fertile pinnae. Spores ellipsoid, with prominent tubercles on distal face. x = 37. Six species: Africa, Asia, Australia, Pacific islands; one species in China. Stenochlaena is exceptional because of its distinct stele type. Ching (1978) gave it family status and placed it after the Acrostichaceae. Kramer (1990) treated it as a subfamily within the Blechnaceae based on its venation, stomata, spores, and gametophyte, which agree best with the Blechnaceae. Recent molecular data show that Stenochlaena forms a clade with species such as Blechnum indicum N. Burman and with Salpichlaena.
1. Stenochlaena palustris (N. L. Burman) Beddome, Suppl. ca. 1 mm in diam., caducous. Fronds distant, dimorphic. Stipe Ferns S. Ind. 26. 1876 [“palustre”]. 10–80 cm, glabrous; lamina 1-pinnate, oblong in outline, 50–80 × 20–30 cm; lateral pinnae 8–16 pairs, terminal pinna similar; 光叶藤蕨 guang ye teng jue pinnae very shortly stalked, articulate to rachis; sterile pinnae Polypodium palustre N. L. Burman, Fl. Indica, 234. 1768; broadly lanceolate or oblong-lanceolate, (10–)15–20 × 2–4.5 Acrostichum palustre (N. L. Burman) C. B. Clarke; A. scandens cm, varying much in size and shape, leathery or stiffly papery, Hooker (1864), not Bory ex Fée (1845); Chrysodium palustre surface smooth and glossy, base broadly cuneate, margin (N. L. Burman) Luerssen; Lomaria scandens Willdenow; sharply and irregularly serrate, apex acuminate or sometimes Lomariopsis palustris (N. L. Burman) Kuhn; L. scandens Met- caudate; fertile pinnae 10–20 × 0.1–0.5 cm, linear. Veins simple tenius; Olfersia scandens C. Presl; Onoclea scandens Swartz, or forked, forming single row of narrow costal areoles. Sori nom. illeg. superfl.; Pteris scandens (Willdenow) Roxburgh; acrostichoid, covering whole abaxial surface of fertile pinnae. Stenochlaena hainanensis Ching & P. S. Chiu; S. scandens J. 2n = 148. Smith. Secondary forests, open places; near sea level to 400 m. Guang- Stems long creeping and climbing; scales at apex of rhi- dong, Guangxi, Hainan, Yunnan [Cambodia, India, Indonesia, Laos, zome dark brown to black, orbicular, ovate, or ovate-lanceolate, Malaysia, Nepal, Thailand, Vietnam; Australia, Pacific islands].